C&S in Pastures Chap 10 22/10/01 1:39 pm Page 193
10 Competition and Environmental Stress in
Temperate Grasslands
Duane A. Peltzer* and Scott D. Wilson
Department of Biology, University of Regina, Regina, Saskatchewan, Canada
Introduction
There is abundant evidence that competition helps
to determine the species composition of temperate
grasslands (Harper, 1977; Lauenroth and Aguilera,
1998; Wilson, 1998, 1999). One approach to
organizing knowledge about competition is to
examine how it varies along environmental
gradients. Here we review which traits confer
competitive ability to plants and the role of compe-
tition along environmental gradients of stress.
Plant competitive ability can be divided into two
components: competitive response and competitive
effect (Goldberg, 1990). Good response competitors
are those species able to resist suppression by competi-
tors. On the other hand, good effect competitors are
able to reduce the performance of other species. Traits
conferring either aspect of competitive ability,
response or effect, may change along stress gradients.
For example, plants from unproductive, stressful envi-
ronments tend to have high below-ground biomass
allocation, long-lived tissues, high nutrient retention
and high tolerance of water and nutrient stress
(Chapin, 1980; Chapin et al., 1993); these traits may
allow them to displace plants from more productive
environments (see review in Goldberg, 1990).
Both field and garden experiments show that
stress-tolerant species can dominate vegetation over
the long term, due to higher levels of nutrient
retention and tissue longevity (Berendse and Aerts,
*Present address: Landcare Research, Lincoln, New Zealand.
© CAB International 2001. Competition and Succession in Pastures
(eds P.G. Tow and A. Lazenby)
1987; Aerts and van der Peijl, 1993; Berendse,
1994). It is not clear, however, whether these traits
identify good stress tolerators or good competitors
in stressful environments (Grime, 1977; Huston
and Smith, 1987; Tilman, 1988; Berendse and
Elberse, 1990). Whereas many studies have examined
competitive interactions along natural and experi-
mental productivity gradients in grasslands (Wilson
and Tilman, 1991, 1993, 1995; Reader and Bonser,
1993; Reader et al., 1994; Peltzer et al., 1998; see also
reviews by Goldberg and Barton, 1992; Gurevitch et
al., 1992), relatively few studies have explored how
suites of traits responsible for stress tolerance affect
competitive ability, or which traits should be related
to competitive ability along environmental gradients.
How does the role of competition vary along gra-
dients of environmental stress? Here we follow
Grime’s (1979) definition of stress as any factor that
limits biomass production. In grasslands, biomass is
typically limited by soil resources: water (Lauenroth
et al., 1978; Sala et al., 1988; Silvertown et al., 1994)
or nutrients (Tilman, 1987; Berendse, 1994). Peak
standing crop (i.e. above-ground plant biomass) is
frequently used as a measure of stress in herbaceous
grassland communities: low standing crop reflects
high stress and low soil resources. Grime (1979) pre-
dicted that stress should limit plant growth rates and
resource demand. Thus, competition might be rela-
tively unimportant on dry or nutrient-poor soils that
support little standing crop. Competition might
193
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194 D.A. Peltzer and S.D. Wilson
increase in importance as soil resources increase and
standing crop increases. This idea can be traced back
to Darwin (Harper, 1977) and is still current
(Keddy, 1989). Alternatively, stressful habitats may
be characterized by intense competition for the
resources that limit growth, such as nutrients and
water (Newman, 1973; Tilman, 1988). As resources
increase, stress decreases, standing crop increases,
shade increases and competition may shift from
being mostly below ground to mostly above ground.
We review evidence for the hypothesis that the
role of competition in controlling community
structure changes along gradients of stress, resource
availability and standing crop. Our focus is on field
experiments that examine the role of competition
and variation in competitive ability along gradients
of stress in natural temperate grasslands. We
include mid-latitude arid and alpine grasslands but
not tropical grasslands or savannahs or Arctic
graminoid communities.
Competitive Ability
The relationship between competitive ability and the
distribution or abundance of species in communities
is still poorly understood, despite many experiments
on competitive interactions in the field (see reviews
by Goldberg and Barton, 1992; Gurevitch et al.,
1992; Goldberg, 1996). Several important questions
remain. What is competitive ability? Does competi-
tive ability vary among species or environments?
How important is competitive ability relative to abi-
otic factors, other ecological processes, and historical
factors (Givnish, 1986; Welden and Slauson, 1986;
Felsenstein, 1988; Keddy, 1989; Harvey and Pagel,
1991; Aarssen, 1992; Underwood and Petraitis,
1993; Westoby et al., 1995; Silvertown and Dodd,
1996)? Much interest has focused on whether the
abundance and distribution of species are the result
of variation in stress tolerance, competitive ability or
niche differentiation among species (Grime, 1977;
Grubb, 1977; Chapin, 1980; Keddy, 1989; Smith
and Huston, 1989; Austin, 1990; Aarssen, 1992;
McLellan et al., 1997). Here we discuss the relation-
ships among competitive ability, stress tolerance and
community composition.
Components of competitive ability
Goldberg (1990) distinguishes between the ability of
plants to perform well in the presence of neighbours
(competitive response), and the ability of plants to
reduce the performance of other species (competitive
effect). Traits conferring success for both response
and effect may change along stress gradients.
Competitive responses
Competitive response ability (sensu Goldberg,
1990) is the ability of a plant to resist suppression
by neighbours (i.e. the resident vegetation). This
can be measured as the change in performance of a
target plant or species in response to the presence of
neighbours, either conspecific or interspecific.
There are several reasons for using small target
plants in existing vegetation (Goldberg, 1990).
First, all plants must regenerate and pass through
critical early life stages in order to establish; early
life stages may be more sensitive to the effects of
neighbours and the environment than established
adult plants (Grubb, 1977). Thus, individual perfor-
mance (e.g. seedling survival and growth) can be
linked with population processes (e.g. recruitment,
population growth, distribution). Secondly, competi-
tive response ability determines which species persist
in a habitat to contribute to community-level diversity
and productivity (Wilson and Tilman, 1995). Thirdly,
good response competitors are able to withstand
resource shortages imposed by competing plants and
are also likely to persist in stressful habitats.
Plant traits associated with competitive response
ability are those which allow a species to persist and
perform well in the presence of neighbours, and
thus resource shortages or stress. Similarly, the best
competitors in Tilman’s (1982) resource ratio
hypothesis model of plant competition are those
species having the lowest resource requirement, or
R*. Empirical support for this model was provided
by Wedin and Tilman’s (1993) experimental mix-
tures of native prairie grasses. In their study, grasses
with the lowest R* values won in competition,
regardless of initial planting densities in mixtures.
Generally, traits suggested to confer competitive
response ability in relatively unproductive systems
are identical to those suggested for stress tolerance
(Table 10.1). Typically, these species have high
root : shoot ratios and low growth rates, are small
and have nutrient-conserving mechanisms, such as
long-lived tissues, carbon-based defences and
storage organs (Grime, 1977, 1979; Chapin, 1980,
1991; Tilman, 1988; Berendse and Elberse, 1990;
Chapin et al., 1990).
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Competition and Environmental Stress 195

Table 10.1. Summary of plant traits suggested to confer high competitive response or competitive effect
ability at low and high primary productivity. The last column shows traits associated with stress tolerance.
Traits for competitive ability are modified from Goldberg (1996).

Response Effect

Low High Low High

productivity productivity productivity productivity Tolerance

Leaf allocation Lowabf Lowaf Low High Lowde

Stem allocation Lowab Higha Low High Lowde
Root allocation Highabf Lowaf High Low Highde
Reproductive allocation Lowab Lowa High Lowcg Lowcdi
Growth rate Lowabf Lowabf High Highc Lowcde
Litter production Low High High Highc Lowc
Photosynthetic rate Lowa High High High Lowe
Height at maturity Lowa Higha High Highc Lowc
Leaf area Low High High Highg Lowci
Plant mass Low Higha High Highc Low
Specific root length Higha High High Highh High
Leaf area/mass Lowaf Highaf Low Highcg Low
Tissue longevity Highag Lowadg High Lowcg Highcdei
Plasticity Low Low High Highc Lowcd
Tissue [N] Lowabg Highbg High High Lowde
Rate of nutrient uptake Lowb Lowb High Highc Lowe
Nutrient storage High Low High Both Highdi
Nutrient losses or leaching Lowb Highb High High Lowd
Nutrient foraging ability High Low High Both Low
Nutrient-based defences Highb Lowb Low Lowc Highe
C-based defences Highe Low Low Low High
Mycorrhizal infection Highg Low Low Low Highd
Shade tolerance Lowbf Highaf Low Lowc Low
Drought tolerance Highf Lowf Low Lowc High
Low-nutrient tolerance Highbg Lowag Low Lowc High

a, Tilman (1988); b, Tilman (1990); c, Grime (1977); d, Chapin (1980); e, Chapin et al. (1993); f, Smith
and Huston (1989); g, Berendse and Elberse (1990); h, Caldwell and Richards (1986); i, Grime (1979).

Why might variation in competitive response be
important in grasslands? Variation in competitive
responses among species may determine species
positions within a competitive hierarchy. This in
turn may determine their distributions along stress
gradients or their relative abundance in a community
(Grime, 1979; Keddy, 1989). The relationship
between species competitive responses and distribu-
tions is relatively well documented in wetlands (e.g.
Wilson and Keddy, 1986; Gaudet and Keddy, 1988;
Keddy and Shipley, 1989) but less well understood
in grasslands (Mitchley and Grubb, 1986; Aarssen,
1988; Wilson and Tilman, 1995). Generally, there is
no strong relationship between competitive
responses and the distribution or abundance of
species along stress gradients (Herben and Krahulec,
1990; Silvertown and Dale, 1991; Shipley and
Keddy, 1993). For example, Wilson and Tilman
(1995) measured the competitive responses of eight
grassland species (four forbs and four grasses) in an
old field in Minnesota. The species showing the
weakest response, i.e. the species most suppressed by
neighbours, was the numerical dominant of natural
vegetation, the perennial grass Schizachyrium scopar-
ium. This result was robust across four combinations
of nitrogen (N) availability and soil disturbance.
STRESS TOLERANCE AND COMPETITIVE RESPONSES.
Because competition is often for resources and
many of the traits associated with stress tolerance
interact with patterns of resource availability, there
should be a close link between competitive response
ability and stress tolerance. This is not a new idea;
Grime (1977) listed traits associated with stress tol-
erators and competitive species two decades ago. For
Grime (1977), any energy a plant spends coping
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196 D.A. Peltzer and S.D. Wilson
with stress decreases competitive ability, so that
there is a trade-off between stress tolerance and
competitive ability. In contrast, other authors define
a good competitor as a species able to perform well
despite resource shortages (Tilman, 1982, 1988;
Berendse and Elberse, 1990; reviewed by Grace
1990, 1991).
The distinction is this: species in productive
environments are competitive if they exploit
resources as quickly and efficiently as possible
(Grime, 1977). Resource competition may also be
intense in unproductive habitats (Wilson and
Tilman, 1991, 1993, 1995; Wilson, 1993a, b), but
the species that lower limiting resources to the
lowest level and use them most efficiently may
dominate the vegetation (Tilman, 1982). Thus,
there may be no trade-off between stress tolerance
and competitive ability. Stress tolerance and com-
petitive ability can be conferred by the same traits
(Tilman, 1982). Traits predicted to confer both
competitive response ability and stress tolerance are
summarized in Table 10.1.
WITHIN-SPECIES VARIATION IN COMPETITIVE
RESPONSES. Within-species or within-population
variability in competitive ability may also con-
tribute to species coexistence and community
composition. Within-species variability may dimin-
ish among-species differences in competitive ability.
Genotypic variability and specificity of interactions
with neighbouring plants or among sites have been
shown in pastures and old fields (Aarssen, 1988;
Mehrhoff and Turkington, 1990; Turkington,
1991). For example, Mehrhoff and Turkington
(1990) compared the competitive ability of five
populations of Trifolium repens from different-aged
pastures. They found considerable variation in
competitive ability among populations of T. repens
in pot, garden and pasture competition experi-
ments with grasses.
Aarssen (1992) reviewed several theories
regarding variation in competitive abilities at the
genotype level; his working hypothesis is that varia-
tion in competitive ability among genotypes within
a species is as great as variation among species. One
implication of this hypothesis is that competitive
exclusion will not occur at the level of the species in
a community. Cheplick (1997) explored within-
species variation in competitive response using 11
genotypes of the rhizomatous perennial grass
Amphibromus scabrivalis grown in competition with
Lolium perenne. Genotypes with more widely
spaced ramets (‘guerilla’ strategy) were more
strongly suppressed by L. perenne than genotypes
producing few, closely spaced ramets (‘phalanx’
strategy). The role of within-species variation in
competitive response in determining species per-
sistence in natural communities deserves further
attention.
RESPONSES TO HETEROGENEITY. Species may respond
differently to resource heterogeneity. Such hetero-
geneity may occur in both space (Campbell and
Grime, 1989; Campbell et al., 1991; Caldwell and
Pearcy, 1994; Casper and Cahill, 1998) and time
(Fitter, 1986; Campbell and Grime, 1989;
Bilbrough and Caldwell, 1997; Goldberg and
Novoplansky, 1997).
Recent studies have shown that plants respond
to soil nutrient patchiness independently from
resource level (Hutchings, 1988; Casper and Cahill,
1998). There is abundant evidence that the ability
to forage for patchy resources differs among species
of grasses (Jackson et al., 1990; Humphrey and
Pyke, 1997; Reynolds et al., 1997). Further, there
may be a trade-off between the precision of
foraging and the size of resource patches exploited
(Campbell et al., 1991). Large, rhizomatous grasses
should be better foragers than tussock grasses, but
their foraging precision should be low, due to wider
spacing between ramets.
Species may also differ in their responses to
temporal heterogeneity (Grime, 1979; Chapin,
1980; Goldberg and Novoplansky, 1997). In nat-
ural grasslands, such as the North American Great
Plains, soil resources and water may be available
only during infrequent pulses (Sala and Lauenroth,
1985; Sala et al., 1992), and differences in ability
to exploit temporal patches may contribute to
competitive success (Caldwell, 1994). Campbell
and Grime (1989) showed that a relatively slow-
growing grass, Festuca ovina, was able to use short
nutrient pulses of from 0.1 to 10 h in duration,
while a faster-growing grass, Arrhenatherum elatius,
could use only longer pulses. Grime (1994) sug-
gests that species occurring in habitats with ‘chronic
nutrient stress’ have large, long-lived root systems,
which remain functional throughout the year and
are capable of utilizing short resource pulses. In
contrast, faster-growing species tend to produce
new tissues to capture additional resources, making
their responses to pulses much slower. Although
resource allocation has received much attention,
grasses may also vary in their physiological ability
C&S in Pastures Chap 10 22/10/01 1:39 pm Page 197
Competition and Environmental Stress
to capture nutrients in response to resource pulses
(Mouat, 1983; Caldwell, 1994). Interest in the
importance of plant responses to resource hetero-
geneity is increasing (Shorrocks and Swingland,
1990; Bell and Lechowicz, 1994; Caldwell and
Pearcy, 1994; Grime, 1994; Miller et al., 1995;
Bilbrough and Caldwell, 1997; Casper and Cahill,
1998), but much remains to be done in grasslands.
Competitive effects
Competitive effect ability (sensu Goldberg, 1990) is
defined as the ability of a neighbouring plant or
species to suppress a focal individual or species.
Competitive effects are quantified as the per-plant
or per-unit-mass (e.g. per-gram) reduction in per-
formance of the focal plant or species (see review by
Goldberg and Scheiner, 1993). Competitive effects
are often assumed to be equivalent among neigh-
bour species, because all plants use the same
resources (e.g. light, water, nutrients) and these
resources are supplied along gradients rather than as
discrete packages (Harper, 1965; Goldberg and
Werner, 1983; Goldberg, 1996). As noted above,
this assumption requires more investigation. Bakker
(1996) found differences in competitive effects
between introduced and native grasses in mixed-
grass prairie. Wedin and Tilman (1993) also found
significant differences between early- and late-
successional grasses; later grasses lowered soil N
availability more than early grasses, which may
explain their eventual dominance of old fields.
Competitive effects may occur at several scales.
In the short term, plants reduce resource levels and
create localized zones of resource depletion
(Caldwell, 1994; Huston and DeAngelis, 1994).
Over longer time periods, plants may modify
nutrient cycles through shoot or root litter quality
or plant–soil community feedbacks; other long-
term feedbacks may arise through mutualisms, such
as mycorrhizas, or through diseases (Allen and
Allen, 1990; Fitter, 1991; Read, 1991; Wilson and
Agnew, 1992; Wedin and Pastor, 1993; Bever,
1994; Stark, 1994; Bever et al., 1997; Kleb and
Wilson, 1997; Wilson, 1998).
EFFECTS ON HETEROGENEITY. Plants can also influ-
ence the spatial heterogeneity of soil resources,
water and light in grasslands (Hook et al., 1991;
Caldwell, 1994). For example, Kleb and Wilson
(1997) used a reciprocal soil transplant experiment
between mixed-grass prairie and aspen (Populus
197
tremuloides) forest. Over one growing season,
prairie grasses significantly lowered the heterogene-
ity of both available N and moisture. Thus, grasses
decreased the patchiness of soil resources.
STRESS AND COMPETITIVE EFFECTS. Stress is the result
of both biotic (due to competitive effects) and abi-
otic (environmental) processes (Goldberg and
Novoplansky, 1997). Interactions between biotic
and abiotic stress may occur if competitive effect
varies with environment. Few studies to date have
separated stress caused by neighbouring vegetation
from stress imposed by the environment. Such stud-
ies are needed to understand the relationship
between biotic and abiotic stresses in grasslands.
Austin (1990) suggests that environmental influ-
ences on species performance are as important as
species interactions. It is clear that both influence
grasses. For example, in an old field in Minnesota,
increasing N availability increased the growth of
transplanted grasses by about 90%; removing neigh-
bours increased growth by 40%; and both increas-
ing N and removing neighbours increased growth
by 150% (see Fig. 1 in Wilson and Tilman, 1995).
Some questions remain. Are species with larger
competitive effects less tolerant of harsh or fluctuat-
ing abiotic conditions? The distinction between
biotic and abiotic stress is important because they
probably act at different scales. Abiotic stress
gradients may occur at larger scales, influencing
entire communities, and filtering the local species
pool by removing those species that are physiologi-
cally incapable of tolerating local environmental
conditions (Harper, 1977). Variation in competi-
tive effect ability, and thus biotic stress, acts to
eliminate species after environmental filtering
through competitive exclusion, similar to
Diamond’s (1975) assembly rules. This model has
been applied to wetlands (Keddy, 1992), but has
not been tested explicitly in grasslands. Part of the
reason may be that there are few contrasting envi-
ronments with discrete boundaries in grasslands;
instead, both biotic and abiotic stresses commonly
occur along gradients.
In summary, good response competitors are able to
withstand resource shortages imposed by neigh-
bours. On the other hand, good effect competitors
are able to strongly suppress the performance of
competing species. However, these two components
of competitive ability are not necessarily correlated
(Goldberg, 1990). One way to compare competitive
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198 D.A. Peltzer and S.D. Wilson
ability among species or along gradients of stress is
by measuring competition intensity, the relative
decline in species performance caused by competi-
tion.
Measuring Competition
in the Field
A simple way to measure the intensity of competi-
tion in the field is to compare the performance of
transplants in plots without neighbours with that in
plots with neighbours. Any difference in perfor-
mance is presumably attributable to the presence of
neighbours. This can be done at sites differing in
some other factor of interest, e.g. stress or grazing
intensity. It can also be done with different trans-
plant species in order to compare their responses.
Comparisons of competition intensity (CI) among
sites or species should be standardized for the effect
of the sites or species identities on transplant per-
formance (Wilson and Keddy, 1986; Grace, 1995;
Miller, 1996) as:
CI = (NN – AN) / NN
where NN is transplant performance (e.g. survivor-
ship, mass, growth, seed production, tiller number)
in plots with no neighbours present and AN is
performance with all neighbours present.
Regression analysis can be used to examine
relationships between CI and stress or standing
crop (e.g. Wilson and Keddy, 1986; Belcher et al.,
1995; Grace, 1995; Miller, 1996; Peltzer et al.,
1998). Care should be taken, however, not to use
transplant performance as a measure of stress: in
this case, the same term (e.g. growth in the absence
of neighbours) ends up on both sides of the regres-
sion equation, producing a spurious correlation.
Alternatively, such experiments can be examined
with analysis of variance (ANOVA) of transplant
performance: a significant interaction between
competition and site treatments would suggests
that competition intensity varies among sites (e.g.
Platenkamp and Foin, 1990; Goldberg
and Scheiner, 1993; Wilson and Tilman, 1995).
ANOVA of performance revealing significant
interactions between transplant species and
competition suggest interspecific differences in
competitive ability; three-way interactions, includ-
ing site, suggest that these differences vary with
stress (Wilson and Tilman, 1995).
Does Competition Vary with
Standing Crop?
Stress is a common phenomenon in natural grass-
lands. Plant productivity, usually measured as
aboveground biomass, is normally limited by soil
resources: water and nutrients (Lauenroth et al.,
1978; Tilman, 1987; Sala et al., 1988; Silvertown et
al., 1994). Stress can occur along natural gradients
– for example, rainfall gradients or north- vs.
south-facing slopes. Stress can also vary through
time with among-year variation in precipitation
and temperature. Here we review how competi-
tion varies along natural and experimental stress
gradients.
Is standing crop a reliable indicator
of stress?
Standing crop should reflect stress from a physio-
logical point of view, but other biotic factors may
influence standing crop, making it a poor indicator
of stress. Standing crop, of course, reflects rates of
biomass removal, as well as production. Thus low
standing crop sites may be produced by high dis-
turbance rates. Standing crop in a Minnesota old-
field grassland was enhanced with three levels of N
addition; controls made a fourth treatment. All N
levels were crossed with four levels of soil distur-
bance applied with a mechanical tiller, which
produced 0, 25, 50 and 100% bare ground at the
start of each growing season. Large mammals and
abiotic disturbance (e.g. fire) were excluded from
the experiment, so that the effects of N availability
and disturbance could be studied independently.
Competition intensity, measured by transplants of
the grass S. scoparium (Wilson and Tilman, 1993)
and, in a later experiment, using transplants of
eight species (Wilson and Tilman, 1995), decreased
significantly with increasing disturbance, but did
not vary with N availability. The results suggest
that low- standing- crop habitats may have low
levels of competition if the low standing crop is
partly attributable to disturbance. Removal of
biomass by disturbance decreases community
demand on resources and decreases competition
(Taylor et al., 1990). In a series of studies, Reader
(1992, 1993) and Reader and Bonser (1993) per-
formed plant removal experiments (i.e. removing
one or more species from intact vegetation (sensu
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Competition and Environmental Stress
Keddy, 1989) at varying levels of stress and her-
bivory. Competition affected transplants only in
plots caged to exclude grazers. In open plots,
grazers were far more important than competition
in controlling plant success.
Natural stress gradients
In natural vegetation, variation in standing crop is
often inferred to reflect variation in stress. Reader et
al. (1994) grew seedlings of Poa pratensis in cleared
plots and intact vegetation in fields in Australia,
Europe and North America. Competition intensity
did not increase with standing crop when all sites
were examined together. Transplant experiments
along a gradient of soil depth and standing crop in
Ontario also found no variation in competition
intensity (Belcher et al., 1995). Similar results were
found in an old field in Michigan (Foster and
Gross, 1997); in this case, however, competition
intensity increased significantly with neighbour-
hood litter mass. A meta-analysis of 34 studies
found no difference in competition intensity
between more stressful habitats (desert and Arctic)
and less stressful habitats (prairies, meadows and
old fields) (Gurevitch et al., 1992).
Other experiments, in contrast, have found
variation in competition intensity. Del Moral
(1983) grew transplants in two alpine grasslands in
Washington. In one neighbourhood, with 50 g
m2 standing crop, neighbours increased transplant
survival. In a second neighbourhood, with 650 g
m2 standing crop, no transplants survived in the
presence of neighbours. This result suggests that
neighbours were facilitative at low standing crop
but competitive at high. Gurevitch (1986) followed
the fate of naturally establishing seedlings of grasses
along a topographic gradient in Arizona.
Neighbours decreased seedling performance more
in moister, low-lying sites than on dry ridge tops.
Reader and Best (1989) removed neighbours in an
old field in Ontario and followed the population
response of the composite Hieracium floribundum
in low- and high-standing-crop sites. Initial
Hieracium densities were made similar by thinning
all plots at the start of the experiment.
Neighbours reduced Hieracium performance at
high-standing-crop sites but had no effect at low
standing crop. Similar results were found for three
species of transplants in the same system (Reader,
1992) and for the grass Poa pratensis (Reader and
199
Bonser, 1993), but not for P. compressa, which was
equally suppressed in both habitats. Bonser and
Reader (1995) used a wider selection of standing-
crop values and found competition intensity to
increase with standing crop. In Montana, grass
neighbours inhibited an annual mustard in a wet
year but facilitated it in a dry year (Greenlee and
Callaway, 1996).
Experimental stress gradients
Other transplant experiments have been performed
along experimental fertility gradients. As in the case
of natural gradients, they produce conflicting
results about variation in competition intensity.
Reader and Best’s (1989) Hieracium experiment
described above found significant variation in com-
petition intensity associated with natural variation
in standing crop, but no variation in competition
intensity was produced by supplying extra water to
the vegetation. Wilson and Shay (1990) removed
neighbours from around established grass tussocks
in Manitoba mixed-grass prairie and found that
neighbour removal increased tussock size to a similar
extent in both fertilized and unfertilized plots.
Burning also had no effect on competition intensity.
Wilson and Tilman (1991) transplanted three grass
species into clearings and intact vegetation within a
5-year-old N addition experiment in Minnesota:
neighbours suppressed transplant growth to the
same extent at all N levels. Similar experiments in a
nearby field with one (Wilson and Tilman, 1993),
two (Wilson, 1994) and eight species (Wilson and
Tilman, 1995) produced the same result. Further,
competition did not increase with added N in
either undisturbed or tilled plots, indicating that
both perennial and annual grass neighbourhoods
had similar behaviour along the experimental fertil-
ity gradient (Wilson and Tilman, 1993, 1995).
DiTommaso and Aarssen (1991) transplanted three
grass species into clearings and intact vegetation in
fertilized and unfertilized plots in an old field in
Ontario and found that fertility had no effect on
competition intensity. Grass transplants grown for
3 years in a 5-year-old N addition experiment in an
old field in Saskatchewan were equally suppressed
at all levels of N availability (Peltzer et al., 1998).
Tree and shrub transplants in the same field were
equally suppressed, regardless of whether the
vegetation was supplied with extra water and N (Li
and Wilson, 1998).
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200 D.A. Peltzer and S.D. Wilson
One exception to the trend of competition
intensity not varying along experimental gradients
is given by Reader (1990), who examined the
impact of neighbours on H. floribundum in
unfertilized and fertilized plots: neighbour removal
increased Hieracium recruitment and survival only
in fertilized plots.
Summary of competition intensity and
standing crop
With one exception, competition intensity did not
vary with stress along experimental gradients.
Competition intensity frequently increased with
standing crop along natural gradients, but not
always. In some cases, the increase in competition
intensity was produced by a single point where
neighbours had no negative effect at low standing
crop (Del Moral, 1983; Bonser and Reader, 1995;
Greenlee and Callaway, 1996). Overall, the results
support the idea that competition intensity
increases with standing crop on natural gradients
more than on experimental gradients (Goldberg
and Barton, 1992).
The results of any experiment depend on the
conditions under which it is carried out, and
measurements of competition intensity have been
carried out within a wide range of standing crop
values (summarized in Belcher et al., 1995). We
summarized the results of several studies that gave
the growth rate of grass transplants with and with-
out herbaceous neighbours. Some studies reported
final transplant mass (Wilson, 1994; Gerry and
Wilson, 1995) but were included because they
reported initial mass and allowed calculation of
growth rates. The studies were performed in fields
abandoned from cultivation for at least 10 years.
We included studies in which both neighbour roots
and shoots were removed and for which standing
crop could be determined (Wilson and Tilman,
1991, 1993, 1995; Wilson, 1993a, 1994; Reader et
al., 1994; Gerry and Wilson, 1995; Bakker, 1996;
Foster and Gross, 1997; Peltzer et al., 1998; S.D.
Wilson, unpublished data; D.A. Peltzer and S.D.
Wilson, unpublished data). We examined grass
transplants with herbaceous neighbours, in order to
avoid drastic differences between transplant and
neighbour morphology (e.g. grasses vs. trees). We
excluded two young sites from Reader et al. (1994)
because the topsoil had been removed and the plots
were undergoing primary succession (H. Olff,
1996, personal communication). Competition
intensity (CI) was calculated as above for each
species, year and vegetation type.
There was no clear relationship between compe-
tition intensity and standing crop (Fig. 10.1), either
for the complete data set or for the data set with the
large study of Reader et al. (1994) excluded. Thus,
our summary of other studies corroborates the lack
of an obvious pattern in competition intensity
found by Reader et al. (1994). Although there is
always a danger of falsely failing to reject the null
hypothesis of no effect, an analysis of the same data
set, but including recently disturbed plots, found
that competition intensity did increase significantly
with field age (Wilson, 1999). This disparity
suggests that standing crop is a relatively poor
predictor of competition intensity. Habitats which
are so stressful that plants cannot maintain live
mass are, of course, likely to have little competition
(e.g. Grubb, 1992; Belcher et al., 1995; Kadmon,
1995; Goldberg and Novoplansky, 1997), but
this description probably does not apply to tem-
perate grasslands.
Lastly, dominant species may differ in their
competitive effects regardless of stress, so that in
any particular environment, the intensity of
competition is determined by the identity of the
dominant species and not by stress or resource
availability. Bakker (1996) grew transplants with
and without neighbours in two sections of a 50-
year-old Saskatchewan field. One section had been
planted with the introduced pasture grass Agropyron
cristatum, and the other section had undergone
natural succession to native prairie grasses.
Competition intensity varied little between
dominant vegetation types, but the F ratio for the
competition term in the ANOVA of transplant
growth was twice as high in the Agropyron-
dominated vegetation than in the native-
dominated vegetation, suggesting that Agropyron
exerted greater competitive effects than did native
species (Underwood and Petraitis, 1993). Further,
the growth of transplants of the native grass
Bouteloua gracilis decreased significantly with neigh-
bour mass only when grown in native vegetation,
and the growth of transplants of Agropyron
decreased significantly with neighbour mass only
when grown in Agropyron-dominated vegetation.
These results suggest that competition was most
intense in intraspecific pairings and that variation
in competitive effects occurred without variation in
abiotic stress. Similar results were found for the
C&S in Pastures Chap 10 22/10/01 1:39 pm Page 201

Competition and Environmental Stress 201

Wilson and Tilman, 1991

Wilson and Tilman, 1993
2
Competition intensity

Wilson, 1993a
Reader et al., 1994
Wilson, 1994
1 2070
Gerry and Wilson, 1995
Wilson and Tilman, 1995
× Bakker, 1996
1205 Foster and Gross, 1997
0
× Peltzer et al., 1998
S.D. Wilson, unpublished data
× D.A. Peltzer and S.D. Wilson,
–1 unpublished data
0 200 400 600 700
Standing crop (g m–2)
Fig. 10.1. Competition intensity as a function of standing crop. Competition intensity was measured as the
relative reduction in grass transplant growth caused by grass neighbours. Standing crop increases with
decreasing stress. Data were taken from several studies in natural temperate grasslands.

grass Anthoxanthum odoratum in California, where
A. odoratum is more suppressed by intraspecific
neighbours on dry soils than by interspecific neigh-
bours on moister soils (Platenkamp and Foin,
1990). Goldberg et al. (1995) outline a method for
testing whether differences in competition intensity
are attributable to differences in stress or simply to
differences among neighbouring species.
Stress and Root Competition
If competition is equally important at all levels of
stress, the mechanism of competition may still shift
from below ground to above ground as stress
decreases and standing crop increases. Field
experiments suggest that competition in grasslands
occurs mostly below ground and that neighbour
shoots have little effect on transplant performance
(Cook, 1985; Snaydon and Howe, 1986; Wilson
and Tilman, 1991, 1993, 1995; Seager et al., 1992;
Wilson, 1993a, b; Belcher et al., 1995; Peltzer et
al., 1998; see review by Casper and Jackson, 1997).
Cook and Ratcliff (1984) used root exclusion tubes
of varying depths to show that the intensity of root
competition decreased as neighbour-free soil vol-
ume increased.
Shoot competition occurs in some grasslands if
standing crop is increased through fertilization.
Transplant growth in unfertilized Minnesota old-field
plots was controlled entirely by root competition but
was influenced by both root and shoot competition
in fertilized plots with less light penetration (Wilson
and Tilman 1991, 1993, 1995). This did not occur,
however, in a grassland with a smaller range of
standing crop (Belcher et al., 1995): in this case,
competition was always below ground, regardless of
neighbour mass. Competition was also entirely below
ground in species-poor experimental plots, in which
fertilization did not produce changes in species
composition (Peltzer et al., 1998). Taken together,
these studies suggest that light competition may
occur only if standing crop is relatively high and if tall
life-forms are available to colonize fertilized plots and
cause shading.
Competitive Ability and Grassland
Community Structure
Variation in competitive ability among species
occurs at the individual level. A next step is to
determine if variation among individual species can
be used to predict their performance in a community
(Goldberg, 1990; Goldberg et al., 1995). Specifically,
does variation in the performance of individuals scale
up to the population and community levels? One
method to assess individual- vs. community-level
competitive ability is the community density series
of Goldberg et al. (1995). This method manipulates
the density of the entire community to levels both
below and above those naturally occurring at a site.
Very low-density treatments are null communities,
C&S in Pastures Chap 10 22/10/01 1:39 pm Page 202
202 D.A. Peltzer and S.D. Wilson
which should have no competition occurring
among plants. As the community density increases,
so do the frequency and importance of species
interactions, including competition. The difference
in performance of species between low- and high-
density treatments is its community-level competitive
ability. For example, species whose relative abundance
increases with community density would have a
higher community-level competitive ability. Other
general attempts to scale plant effects at lower scales
to phenomena at higher scales have been discussed at
length elsewhere (Ehleringer and Field, 1993; Jones
and Lawton, 1995; Bazzaz, 1996).
Plant traits may be used to predict which species
have good competitive abilities, and to test these
predictions empirically (Gaudet and Keddy, 1988;
Keddy, 1989, 1992; van der Werf et al., 1993;
Grime et al., 1997; Rösch et al., 1997; Reader,
1998). A partial list of these traits is given in Table
10.1. Whole communities can be screened for
general patterns of traits associated with abundance
and distribution of species, although this is logisti-
cally difficult. Experimental evaluations of traits for
competitive ability are needed (e.g. Wilson, 1991;
Wedin and Tilman, 1993).
One difficulty with trying to scale up variation
in competitive ability or with using plant traits to
predict competitive ability is that competitive
ability may be inconsistent among sites or habitats
(Underwood and Petraitis, 1993; Miller, 1994;
D.A. Peltzer and S.D. Wilson, unpublished data;
see review by Goldberg, 1996). For example,
Vinton and Burke (1997) examined the effects of
prairie plants on carbon and N cycling at three sites
varying in annual precipitation. They found that
differences among species were greatest at the two
most productive sites. This result was probably due
to patchy plant cover at the driest site (the compari-
son was between bare ground and plant cover) and
a more continuous cover at the two wetter sites (the
comparison was among species). In a review of
studies comparing species’ competitive abilities
between environments, Goldberg (1996) found
that competitive responses tended to be consistent
among environments while competitive effects
often varied among environments. Further work is
needed to examine how species’ competitive
effects vary with environment and why. Factorial
experiments designed to observe species effects 
environment interactions are ideal for determining
variation in competitive abilities among species
(Goldberg and Scheiner, 1993).
Interactions between Grazing
and Competition
Grazers have a number of impacts on native grass-
lands: they can remove 9–57% of net above-ground
foliage production (Frank et al., 1998), promote
shoot growth by removing older, less productive tis-
sue (Caldwell et al., 1981; McNaughton, 1983,
1984), enhance nutrient availability by increasing
rates of nutrient cycling (McNaughton et al., 1989;
Day and Detling, 1990; Holland et al., 1992) and
create small-scale disturbances by trampling or bur-
rowing (Crawley, 1983; Huntley, 1991; Olff and
Ritchie, 1998). In addition, grazers have many
indirect effects in grasslands. For example, removal
of above-ground biomass may enhance primary
production through removal of detritus, increased
soil moisture status and plant water-use efficiency,
reduced fire frequency and altered species composi-
tion (McNaughton, 1984, 1985; Knapp and
Seastedt, 1986; Archer, 1995).
At the level of the individual plant, grazers
reduce survival, growth and fecundity (Crawley,
1983; Bullock, 1996). Several authors suggest that
grazing can promote plant growth through removing
senescing tissue and enhancing subsequent growth,
although the majority of studies do not support
this idea (see review by Belsky, 1986). By damaging
plants, herbivores induce defence mechanisms in
plants such as altered morphology or increased
levels of defence chemicals in leaves (Crawley,
1983; Vicari and Bazely, 1993). Grazing of above-
ground tissues also has consequences below ground.
For example, classic work by Weaver (1950)
showed that intense grazing of above-ground
foliage can strongly reduce below-ground biomass
in short-grass prairie. However, the effects of above-
ground grazing on below-ground productivity vary
among grazing systems and the grazing history of the
site (Belsky, 1986; Milchunas and Lauenroth, 1993).
Plants have several anti-herbivore defences
including high concentrations of silica in leaves,
lower-growing meristems, high levels of fibre and
low levels of protein and a variety of defence
compounds (Vicari and Bazely, 1993). Several plant
traits confer resistance to both herbivory and
environmental stress. For example, high tissue
density, tough, fibrous leaves and high concentra-
tions of secondary compounds confer resistance to
herbivores and stress (Chapin, 1980, 1991; Chapin
et al., 1990; Grubb, 1992; see Table 10.1). In
C&S in Pastures Chap 10 22/10/01 1:39 pm Page 203
Competition and Environmental Stress
addition, many traits allowing grasses to withstand
grazing may actually be adaptations to a semi-arid
environment (Coughenour, 1985).
Grazing may have bigger impacts in stressful
habitats than in more productive habitats, for at
least two reasons. First, fewer nutrients are usually
available for plants to take up and use for regrowth
after biomass removal (Grime, 1979). Secondly,
shoot herbivory usually results in a decline in
critical mycorrhizal mutualists associated with plant
roots (Gehring and Whitham, 1994), further
reducing the ability of grazed plants to take up
nutrients needed for compensatory growth.
Herbivory may alter the outcome of competition
between grassland plant species. More specifically,
grazers can influence plant competitive ability
either by modifying growth and morphology or by
influencing plant abundance and distribution
(Louda et al., 1990). Further, by removing standing
biomass, grazing acts to shift plant competition
from shoots (i.e. for light) to roots (i.e. for soil
nutrients or water) (Milchunas et al., 1992). A
common assertion is that herbivores shift the balance
of competition by preferentially consuming domi-
nant competitors, altering the relative competitive
ability of species and promoting the abundance of
plants that are otherwise rare or excluded (Tansley
and Adamson, 1925; McNaughton, 1985; Huntley
1991; Clay et al., 1993; Olff and Ritchie, 1998).
This view is supported by a number of studies. For
example, selective grazing of the late-successional
grass S. scoparium results in the increase of two
grasses that are normally poorer competitors,
Bothriochloa saccharoides and Stipa leucotricha
(Anderson and Briske, 1995). Grazing in a south-
ern English pasture increased the abundance of the
N-fixing forb T. repens and decreased L. perenne,
because the taller Lolium lost more above-ground
mass to grazers (Parsons et al., 1991). For two
grasses growing in low-nutrient soils, clipping both
caused greater reductions in the growth of A. elatius
than in that of Festuca rubra and accelerated the
replacement of Arrhenatherum by Festuca (Berendse
et al., 1992). Below-ground insect herbivory may
also accelerate succession to a grass-dominated
sward by reducing the size and growth of perennial
forbs and perhaps their competitive ability (Brown
and Gange, 1990, 1992). Thus, herbivore-induced
changes of competitive interactions can drive
species replacement in grasslands.
The argument that selective grazing causes the
competitive replacement of palatable species by
203
unpalatable species assumes that a trade-off should
exist between competitive ability and grazing
tolerance (Moretto and Distel, 1997). For example,
plant height should increase a plant’s competitive
ability for light, but may also increase its proneness
to grazing (Gaudet and Keddy, 1988; Oksanen,
1993). A trade-off between competitive ability and
grazing tolerance should occur for competitive
effect ability, but not necessarily for competitive
response ability; this is because many of the plant
traits conferring competition response or stress
tolerance ability also confer resistance to herbivores
(see Table 10.1).
Grazers may be essential for the restoration of
native grasslands. Fire is often used in prairie
restoration across North America, but repeated
burning can lower plant species diversity (Collins et
al., 1995). Recent work by Collins et al. (1998)
found that both mowing and grazing by bison
enhanced species diversity after fire by selectively
reducing the abundance of dominant C4 (warm-
season) grasses, thus reducing their competitive
effects and allowing subordinate C3 (cool-season)
grasses and forbs to persist and contribute to grass-
land diversity. Restoration of diversity by grazers
may be successful at small spatial scales (i.e.
< 10–100 m2), but recent studies suggest that graz-
ing reduces plant diversity over larger spatial scales
(i.e. > 1000 m2) (Olff and Ritchie, 1998; Stohlgren
et al., 1999a, b).
As discussed above, we know that grazing or
herbivory strongly affects the survival, growth and
reproduction of plants, but how does herbivory
modify the outcome of competition along gradients
of stress (Connell, 1975; Menge and Sutherland,
1987; Louda, 1989; Louda et al., 1990)? Most
studies have examined the effects of grazers on
range condition or species composition, or have
used manual clipping in pot experiments to
examine the effects of grazers on plant competition;
relatively few studies have quantified the effects of
herbivory on plant competition under natural
conditions. Further, the effects of different kinds of
herbivores on plant interactions is essentially
unknown, i.e. what are the effects of large-bodied
ungulates vs. small burrowing mammals vs. insects,
or native grazers vs. cattle in the same grassland
system? For example, even though below-ground
herbivory can greatly reduce plant productivity, few
studies have considered the role of invertebrate
grazers on roots, even though most grassland
production is below ground (Stanton, 1988; Brown
C&S in Pastures Chap 10 22/10/01 1:39 pm Page 204
204 D.A. Peltzer and S.D. Wilson
and Gange, 1990). In short-grass prairie, herbivory
by white grubs, the larvae of June beetles (including
Phyllophaga fimbripes), strongly depresses the abun-
dance of the dominant grass, B. gracilis, for up to
14 years after an insect outbreak (Coffin et al.,
1998). Clearly, further work is required to under-
stand how different kinds of above- and below-
ground herbivory interact with competition to
determine the productivity and species composition
of grasslands.
Non-competitive Interactions
Many plant species can coexist in grasslands despite
having large differences in competitive abilities (e.g.
Mitchley and Grubb, 1986; Silvertown et al.,
1994), suggesting that other factors either prevent
the competitive exclusion of species or promote
their coexistence (Bullock, 1996).
Grasses might avoid competition by exploiting
different niches. If this happens, then competition
might contribute little to grassland patterns
(Leibold, 1995). It is difficult to imagine how
plants can divide up resources into distinct niches,
because all plants use the same resources (Harper,
1977; Tilman, 1982; Goldberg and Werner, 1983).
Several studies have explicitly examined resource
partitioning in plant communities (e.g. Werner and
Platt, 1976; Russell et al., 1985), but are often
inconclusive. For example, Mahdi et al. (1988) did
not find significant niche separation among eight
British grassland species using either six niche
dimensions based on soil nutrient levels and
phenology or ratios of N : P as a test of Tilman’s
(1982) resource ratio hypothesis.
In North American prairies, many species
coexist in relatively few habitat types. In these
grasslands, two niche axes are commonly studied:
water use and phenology. Werner and Platt (1976)
showed niche partitioning for populations of six
goldenrod species (Solidago spp.) along a soil mois-
ture gradient in an Iowa prairie, although the same
species did not show niche partitioning in an old
field in Michigan. Many grassland species display
phenological divergence in growth, which is related
to photosynthetic pathway (Ehleringer and
Monson, 1993). For example, Kemp and Williams
(1980) examined photosynthesis, respiration and
growth in two common prairie grasses, Agropyron
smithii (C3) and B. gracilis (C4). Bouteloua gracilis
grew best in warmer conditions, but both species
responded similarly to water stress. The authors
suggest that niche separation may occur along
seasonal temperature gradients rather than seasonal
moisture gradients for these two grasses. Sala et al.
(1982) found that A. smithii took up soil moisture
earlier in the growing season than did B. gracilis.
Similar results were reported for the same two
grasses by Christie and Detling (1982), using a
replacement series experiment. Similarly, many
authors suggest that the success of the invasive
grass Bromus tectorum in large areas of the western
USA is at least partly due to its growth earlier in
the season than native grasses and shrubs (Mack,
1986). Niche space can also be divided up either
through different regeneration niches (Grubb,
1977) or through ratios of limiting nutrients
(Tilman, 1982).
Because there is no theory predicting which
niche axes species should segregate along, it is
impossible to determine if no niche segregation
occurs among species or whether the wrong niche
axes were measured. Thus, there are difficulties with
interpreting what niche differences mean when
they are found and whether they are the cause or
the result of species coexistence (Arthur, 1982;
Silvertown, 1983). Leibold (1995) linked tradi-
tional theory of the niche with mechanistic models
of community interactions (e.g. Tilman, 1982;
Holt et al., 1994), which may be a more promising
approach to testing niche theory.
More recent alternative explanations of species
coexistence have been developed which do not
invoke niche differentiation. Silvertown and Law
(1987) review some of these, including spatial
aggregation in populations (Yodzis, 1986), non-
transitive competition (Keddy, 1989; Shipley,
1993), a variety of non-equilibrium theories
(Shmida and Ellner, 1984; Chesson and Huntly,
1997) and density-independent mortality (Huston,
1979). For example, Ågren and Fagerström (1984)
demonstrated that two species with similar juvenile
competitive abilities and lifespan can coexist if
seed production (i.e. reproductive events) varies
stochastically. Other models have demonstrated
that spatial patchiness or environmental hetero-
geneity promotes species coexistence, at least for
annual communities (Pacala and Tilman, 1994). In
a recent review, Chesson (1991) suggests that we
look for trade-offs that can determine coexistence
and stability in communities – for example,
dispersal vs. competitive ability in grassland plants
(Tilman, 1997).
C&S in Pastures Chap 10 22/10/01 1:39 pm Page 205

Competition and Environmental Stress 205

Conclusions linked to the abundance or distribution of species.

Field experiments suggest that competition controls
plant performance at all levels of stress in natural
grasslands (see Fig. 10.1). Further, competition is
primarily among roots for below-ground resources,
such as water and nutrients. The relative importance
of shoot competition can increase as stress decreases,
shoot mass increases and light penetration decreases.
In addition, results from previous studies indicate
that competition intensity changes more along
natural stress gradients than along experimental
gradients: we need studies of the differences
between natural and experimental stress gradients
in the same system. Particularly for natural gradi-
ents, we need to separate the effects of abiotic and
biotic stress, in order to determine the relative
role of environment and species interactions in
controlling the species composition in grassland
vegetation.
Using plant traits to predict individual- and
community-level competitive ability is a powerful
approach to testing current competition theory.
Screening experiments using several species in
environments differing in stress or productivity are
needed; interactions among species, competition
and environment should be examined to determine
how competitive ability changes with abiotic and
biotic stresses. Results to date indicate that many of
the traits that characterize stress-tolerant species
should also enable such species to be effective
competitors in stressful habitats.
Field experiments have failed to show that an
ability to perform well in the presence of neigh-
bours (i.e. good competitive response ability) is
In contrast, grasses that are best able to reduce
resources (i.e. good effect competitors) tend to
dominate nutrient-poor soils, suggesting that com-
petitive effect ability may determine the relative
abundance of species in grasslands.
In the process of resource consumption, grasses
may also affect the patchiness of resources. The
scale of this patchiness is in turn likely to influence
other plants. Our understanding of the relative
contributions of competition and stress to pasture
community composition will be enhanced by closer
study of the dynamics and heterogeneity of below-
ground processes.
Understanding how competitive ability and
environment interact is increasingly important
because grassland environments are often altered by
humans through fertilization, nutrient pollution,
mowing, grazing and the introduction of new
species. Moderate levels of grazing can interact
with plant competition to maintain the diversity
of grasslands and offset declines in plant diversity
caused by frequent fires. Thus, grazing may be an
important tool for the restoration of grassland
diversity. However, more work is needed on
comparing the effects of different grazers on the
outcome of competition in the same system.
Acknowledgements
We thank the Natural Sciences and Engineering
Research Council of Canada for support and
P. Tow and A. Lazenby for helpful comments on an
earlier draft of the manuscript.

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